The Effects of Size and Habitat on Delta N-15 of Carnivorous Plants
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.I THE EFFECTS OF SIZE AND HABITAT ON 8 N-15 OF CARNIVOROUS PLANTS (Drosera spp.) supervisors: Dr. J. J. Midgley Prof. w. D. stock Project by : Theresa Nobuhle Mgidi HONOURS PROJECT 1999 University of Cape Town The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University of Cape Town BOLUS LIBRARY C24 0007 6233 CONTENTS 1111111111111~~ Page# Abstract 1. INTRODUCTION 1 1.1) Definition 1 1.2.1) Evolution of carnivory 1 1.2.2) Alternatives for the evolution of carnivory 3 1.3) Aims 4 1.4) Predictions or Hypotheses 6 2. MATERIALS AND METHODS 8 2.1) Materials 8 2.2) Study sites 9 2.3) 8 N-15 Natural Abundance Measurement 10 _, 2.4) Measurements of Chlorophyll and Anthocyanins 11 2.4.1) Chlorophyll 11 .,r 2.4.2) Anthocyanins 12 ,_\ \_ j LJ . } 3. RESULTS 13 ' 3.1) 8 N-15 Measurements 13 ( \ 3.2) Chlorophyll and Anthocyanins 15 ' 4. DISCUSSION 17 4.1) Sun vs Shade 17 4.2) Tall vs Short Drosera plants 20 4.3) Cool/Damp vs Warm/Seasonally Wet 20 4.4) Chlorophyll and Anthocyanins 21 5. CONCLUSION 23 6. ACKNOWLEDGEMENTS 24 7. REFERENCES 25 Theresa Nobuhle MgidiO 0 Department of Botany, University of Cape Town ABSTRACT: The 8 N-15 natural abundance method was qsed to investigate the role of nutrient poor habitats in carnivorous Drosera capensis and Drosera aliciae, and how that role changes under sunny and shady environmental conditions. The main purpose of the study was to evaluate Givnish's (1984) cost/benefit model used to explain the evolution of carnivory in nutrient-poor, sunny and moist habitats. 8 N-15and total nitrogen values of the Drosera species were compared against each other, as well as against the non-carnivorous reference plants collected from each of the two habitats. Generally, data indicated significant differences between the carnivorous plants and their reference plants in terms of 8 N-15 values. However, there was no significant difference between plants collected from the shade and those collected from the sun for both Drosera species. Total nitrogen results revealed higher values for Drosera plants from Camp's Bay than those from Table Mountain did. This indicated that there was a bigger source of insect nitrogen at that site, meaning more insects were available and being caught by the plants at Camp's Bay. Further investigations were performed on the two Drosera species in order to find the influence of altitude, leaf size and plant form, on the degree of carnivory. There was an overall, higher degree of carnivory at Camp's Bay where it is, seasonally wet and the plants have longer leaves and a stem-like rosette. On Table Mountain it is cooler, waterlogged, and the plants have short leaves and ground-level rosettes therefore, the degree of carnivory there was lower. Lastly, chlorophyll and anthocyanin contents were measured and compared between the sun and shade collected D. capensis plants, with tentacles intact and with them removed. Chlorophyll investigations showed significant differences between sun and shade collected D. capensis plants but these were not affected by the removal of tentacles. Alternatively, anthocyanin measurements indicated that sun and shade collected D. capensis plants have similar amounts of anthocyanins, but the removal of tentacles results in a decrease (about four times lower) in the anthocyanin content. 1 1] INTRODUCTION 1.1) Carnivory defined: According to Givnish (1984) the definition of plant carnivory is composed oftwo distinguishing characteristics: an ability to attract, capture and digest prey; and an adaptation to absorb nutrients from prey using the plant's surface, thereby increasing the plant's fitness (i.e. increased growth, survival and reproduction). 1.2.1) Evolution of carnivory: Carnivorous plants are usually found in nutrient-poor environments (Darwin, 1875). However, when Givnish (1984) studied an epiphytic Bromiliad, Brocchinia reducta, which also occurs in nutrient limited environments, he noticed that the Bromeliad was not carnivorous and that few epiphytes were carnivorous. The epiphytes were found in nutrient-poor environments that were either shady or exposed to sunlight and regular desiccation. In contrast, carnivorous plants were found in sunny, moist (at least seasonally) nutrient-poor habitats. Similar observations (Darwin, 1875)on the conditions in which carnivorous plants were found had been published prior to Givnish (1984) but no explanations for the preferences were given. Givnish (1984) explained the carnivory in plants using a cost/benefit model. The model considers the energetic benefits and costs of carnivory in various habitats. It is based on the theory that carnivory should evolve if the benefits are greater than the cost of investments of adaptations to carnivory, so that the plants with the mutations for such investments should have an advantage when competing with other plants (Givnish 1979, 1982). The cost/benefit model of plant carnivory is made up ofthree potential benefits and these all support the preference of sunny, moist nutrient-poor habitats by such plants. The primary benefit of this model is the enhancement of photosynthesis due to an increase in nutrient absorption and consequently increased growth and reproduction rates. The enhancement of photosynthesis by the increased nutrient absorption rate depends on environmental conditions (Givnish, 1984). As the amount of energy for carnivory (e.g. traps and/or digestion enzymes) increases, the amount of nutrients absorbed should also increase. Furthermore, as the amount of energy for carnivory \ 2 i i continues to increase, the photosynthetic benefits should rapidly increase and then ' slowly level-off as an equilibrium is reached. 'f.his indicates that factors other than nutrients limit photosynthesis and hence, carnivory in plants. The effective rate of photosynthesis is unlikely to increase if nutrient availability increases, unless nutrients are in short supply and limit photosynthesis (Givnish, 1984). For example, studies by Soreson and Jackson (1968) on a Utricularia species and Chandler and Anderson (1976) on a Drosera species showed that usual increase in growth of carnivorous plants supplied with prey on nutrient-poor habitats largely disappears as nutrient availability in the habitat increases. Therefore, the greatest benefit is expected in nutrient-poor sites (Givnish, 1984). However, if factors like light or water are limiting then the extent to which nutrients added by carnivory can increase photosynthesis decreases (Bannister, 1976). A study by Gulmon and Chu (1981) showed that photosynthesis increases more slowly with added leaf nitrogen at low light intensities than at high ones. Thus, sunny environments are more favourable for maximum benefits from carnivory (i.e. photosynthesis) than shady habitats. f- 0: 0 a: a.. !,2 f UJ I fz (/)> ~ it PHOTOSYNTHETIC COST I INEFFICIENCY ASSOCIATED WITH CARNIVORY Figurel: Photosynthetic benefits and costs associated with differential levels of investment in carnivory adaptations in nutrient-poor sites, as a function of environmental conditions [Taken from Givnish, 1984] Figure 1 shows that the benefits of carnivory and consequently, photosynthesis, are limited by the availability of water and light. The relationship is such that in habitats that are sunny and moist carnivory yields greater benefits than in shady and/or dry habitats. Overall, Givnish' s cost/benefit model suggests that the differences in ph<?tosynthetic benefits and costs are positive (i.e. benefits> costs) when there is not so much investment in carnivory. This happens in nutrient-poor environments that are also sunny and moist. 3 1.2.2) Alternatives for the evolution of carnivory: Zamora (1999) has suggested other reasons for carnivory being abundant in sunny and moist, nutrient-poor habitats, besides the cost/benefit reasons put forward by Givnish (1984). Zamora (1999) does not dispute Givnish's (1984) theory; he adds dimension to the subject by giving alternative reasons of carnivory dominating in sunny environments. He looked at the bladders in an Utricularia species, which are used by the plant to trap insects, and noted that the number ofbladders increased in nutrient rich soils. This result is opposite to expectations suggested by Givnish (1984). Therefore, according to Zamora (1999) the degree of carnivory is higher in nutrient rich environments than nutrient limited ones. Bronstein (1994) showed that results of interactions between organisms are affected by current ecological conditions. Factors such as size and age of carnivorous plants as well as density and population structures of insects can determine the nature of plant-insect interactions. This is such that older and larger leaves have the capacity and opportunity to catch more insects and areas abundant in insects increase the likelihood of insects being trapped by carnivorous plants. Also, abiotic conditions can play a major role in determining the outcome of such plant-insect interactions (Dunson and Travis, 1991) because both plants and insects are affected by environmental conditions, these conditions either promoting the survival of plants and insects or leading to a decline in their numbers. This latter concept supports Givnish' s reasons for carnivory being most successful in sunny, moist environments. The site-specific effects (i.e. shady, sunny) in plant-insect relationships are particularly important because the environment governs the ecophysiology of both plants due to their sessile lifestyle, and insects due to their small size (Zamora, 1999). With all these points in mind, reasons for sunny nutrient-poor habitats being favoured by carnivorous plants, may also be due to the greater abundance of insects of all sizes in open sunny habitats than in shady and dry ones.